lepidopteran (order Lepidoptera), any of more than 155,000 species of butterflies, moths, and skippers. This order of insects is second in size only to Coleoptera, the beetles.

Because of their day-flying habits and bright colours, the butterflies are more familiar than the chiefly night-flying and dull-coloured moths, but the latter are far more varied and abundant. The skippers are a worldwide group intermediate between butterflies and moths. With the exception of a few moths, all adult lepidopterans have two pairs of wings. The name Lepidoptera is derived from the Greek, meaning “scaly winged,” and refers to the characteristic covering of microscopic dustlike scales on the wings.

The life cycle of lepidopterans consists of four stages: egg, larva (caterpillar), pupa (chrysalis), and adult (imago). The larvae do most of the eating, with the majority feeding on foliage, although many species eat stems, roots, fruits, or flowers. A number of moth and a few butterfly larvae are serious pests in agriculture and forestry. The adults of many species are important for their role in pollination, which occurs as they visit flowers for nectar. In most land environments the lepidopterans are ecologically important because they transform large amounts of plant matter into animal matter and in turn serve as food for many other groups of animals.

Many members of the order, especially the butterflies, have appealed to the human imagination for thousands of years as symbols of fragile and ephemeral beauty. References to them abound in literature, and they have been depicted in many paintings, have inspired the designs of jewelry, ornaments, and textiles, and have even occurred in many heraldic devices and on postage stamps.

Size range and distribution

Moths, butterflies, and skippers show great diversity in size and development rates. Some moths have wingspans as small as 4 mm (0.13 inch), whereas the largest moths and butterflies measure nearly 30 cm (about 1 foot). Fast-developing species may complete their development in as little as three weeks, while slower ones may require as long as two or even three years.

Lepidopterans live on every continent except Antarctica. Though they are far more numerous and diversified in the tropics, some species survive at the limits of polar vegetation. There are many successful species in nearly every environment, from arid deserts and high mountaintops to marshes and tropical rainforests. Most have become adapted for life in relatively restricted ecological niches and may be limited to only one or a small group of food plants, often to only a single part of one plant. They are therefore seldom abundant in more than one type of habitat. However, a few species have broader food habits and may occasionally reach peaks of abnormal abundance and defoliate large areas of deciduous forest or grassland. Some of the insects most injurious to human interests in agriculture and forestry are members of the Lepidoptera.

The large family-level groups and many smaller ones are worldwide in distribution, with owlet moths (family Noctuidae), measuring worm moths (family Geometridae), pyralid, or snout, moths (family Pyralidae), brush-footed butterflies (family Nymphalidae), and skippers (family Hesperiidae) being dominant elements of the insect fauna everywhere. A few families are characteristically more abundant in one faunal region than in others. Central and South America are characterized by great diversity of American false tiger moths (family Notodonidae) and ctenuchid moths (family Arctiidae). The North American (Nearctic) and Eurasian (Palaearctic) families show much evidence of close connections, chiefly between Asia and western North America. Each region has, however, many distinctive genera.

Many lepidopterans exist only in isolated colonies as relict (remnant) populations, cut off from relatives elsewhere by geologic or climatic changes. Australia and New Zealand have unusually diverse relict populations of the primitive mandibulate moths (family Micropterigidae) and swifts, or ghost moths (family Hepialidae). In North America, Europe, and Asia, many relict species have survived since the last ice age on isolated southern mountaintops.

Importance

Many hundreds of Lepidoptera injure plants useful to humans, including some of the most important sources of food, fabrics, fodder, and timber. The great majority of the injurious species are moths, and the detrimental life stage is always the larva. However, unlike members of other insect orders, lepidopterans do not act as carriers of plant diseases, nor are any of them parasites of or injurious to humans. However, some species feed on open wounds or bodily secretions of wild or domestic animals.

The list of valuable plants subject to damage by lepidopterans is a long one, including many grains, sugar beets and sugarcane, cotton, tobacco, some root crops and leaf crops, many fruits, and timber and shade trees. The damage may involve the leaves, stems, roots, or fruit. Woolens, furs, silk, and even feathers are eaten by fungus moths (seetineid moths) of several genera (clothes moths). The greater wax moth (Galleria mellonella) causes considerable damage in beehives.

A few Lepidoptera are directly beneficial to humans. Nearly all silk is obtained from the domesticated silkworm (Bombyx mori), which is originally from China. Other silks such as shantung and tussah are the products of various Asiatic giant silkworm moths (family Saturniidae). The larvae and sometimes the adults of a few species are used for food. The larvae of one skipper (Rhopalocampta libeon) are collected in large quantities in the Congo, and the 10-cm (4-inch) caterpillars of giant skippers (family Megathymidae), known in Mexico as gusanos de magüey, are both consumed domestically and canned and exported for consumption as hors d’oeuvres. The South American cactus moth (Cactoblastis cactorum) has been highly beneficial in weed control, clearing more than 150 million ha (60 million acres) in Australia of alien prickly pear cactus. Doubtless, humans also benefit from much unrecognized weed eating by caterpillars and flower pollination by adults.

Many lepidopterans are valuable in biological research, including work in ecology, biogeography, systematics, genetics, and physiology. Much of the present knowledge of endocrine controls of insect development has come from studies of the silkworm moth and its relatives. The study of the British peppered moth (Biston betularia) has profoundly influenced ideas about rates of evolutionary change. An increase in the proportion of dark moths, a change thought to be brought on by airborne soot produced during the Industrial Revolution, has been termed industrial melanism.

Natural history

Feeding habits

The habits of Lepidoptera are extremely diverse, depending on the adaptations of the species or group to climate, environment, type of food plant, way of feeding, and many other factors. The great majority of food plants are conifers and flowering plants, but primitive plants such as mosses, liverworts and ferns, and some lichens are eaten by a few groups. Nearly all parts of the plant are eaten by various specially adapted caterpillars. Flowers are eaten by many larvae, including plume moths (family Pterophoridae), with nectar being consumed by many adults. Cones and fruits and their seeds are eaten by others, such as yucca moths (family Incurvariidae), leaf roller moths (family Tortricidae), and some owlet moths. Some seed eaters such as the flour moth (genus Ephestia) have become household pests, feeding on stored grains and cereals. Buds or soft, succulent stems are bored into by members of many families. Several lepidopteran groups—for example, the pine moth (Rhyacionia)—specialize on the terminal shoots of conifers. Various groups feed on the turf of grasses and sedges. The carpenter (family Cossidae), ghost (family Hepialidae), and clearwing moths (family Sesiidae) bore into woody stems and rootstocks. Carpenter moths in particular tunnel deep into hardwoods. A good many lepidopterans, especially fungus moths (family Tineidae), scavenger moths (family Blastobasidae), and snout moths (family Pyralidae), feed on dead and decaying plant matter, particularly moldy debris. In comparison with other orders of insects, relatively few Lepidoptera live in plant galls or eat animal matter. Several families contain leaf miners, especially Gracillariidae (leaf miner moths), Elachistidae (grass miner moths), and Tischeriidae (trumpet leaf miner moths).

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